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On the So-Called Pharyngeal Gland-Cells of Earthworms. by J

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On the So-Called Pharyngeal Gland-Cells of Earthworms. by J PHARYNOEAL GLAND-CELLS OF EARTHWORMS. 253 On the So-called Pharyngeal Gland-cells of Earthworms. By J. Stephenson, D.Sc, M.B., Lieutenant-Colonel, Indian Medical Service; Professor of Zoology, Government College, Lahore. With Plate 19. CONTENTS. PAGE 1. HISTORICAL ...... 253 2. MATERIAL AND METHODS .... 260 3. PHEEETIMA POSTHUMA .... 261 4. PHERETIMA HETEROCH,S:TA , 265 5. PHERETIMA HAWAYANA .... 267 6. LUMBRICID.*: . ... 269 7. THE APPEARANCES IN YOUNG SPECIMENS . 274 8. THE CELLS IN THE LUMBRICID EMBRYO . , 279 9. FUNCTION OF THE CELLS . .'281 10. SUMMARY . .283 11. REFERENCES TO LITERATURE .... 284 12. EXPLANATION OF FIGURES .... 2S5 HISTOIUCAL. SUCCEEDING the buccal cavity in all earthworms is a swollen portion of the alimentary tube, the pharynx. The usual description of this portion of the tube in Lumbricus may be given in the words of Parker and Haswell (7): the "buccal cavity ... is followel by a much larger thick-walled, rounded chamber, the pharynx. From the wall VOL. 62, PART 3. NEW SERIES. 19 254 J. STEPJdENSON. of the pharynx there run outwards to the body-wall a number of radially arranged bundles of muscular fibres which, when they contract, draw the pharynx backwards, and at the same time dilate it." One of the constituents of this pharyngeal thickening, not mentioned in the ordinary descriptions of the earthworm, is nevertheless a prominent feature, easily visible under tlie lens in the ordinary dissection, and immediately obvious, owing to its staining properties, in sections through the region where it occurs. This constituent is a cellular mass which forms soft white projecting lobules on the dorsal and lateral aspects of the pharynx; the lobules surround the muscular strands which issue from the pharynx, and in addition, the cells of the mass penetrate inwards between the interlacing muscular bundles of the thick dorsal pharyn- geal wall in the direction of the lumen of the canal. Though these cells have received some attention from previous writers, an adequate account of their nature and origin has not yet, I believe, been given. References to previous authors are given by Vejdovsky (9, 1884), from whose account of them I quote, since the older literature is inaccessible to me. The earlier investi- gators—Leo, Clarke, Lankester—who saw these masses of pharyngeal cells in Lumbricus, interpreted them as glandular. Perrier described pharyngeal glands in several genera; in Pontodrilus they are said to be variously coiled tubes whose walls are composed of large cells with granular contents; in Moniligaster they pour their secretion into the pharynx by a multitude of small, canals visible with the lens; Perichasta houlleti has several layers of glands which open into the interior by three pairs of orifices. Claparede refers to those cells of the pharyngeal mass which penetrate inwards between the muscular bundles as " . numerous polygonal cells with large round nuclei 6 n in diameter. The import of these cells is at present not clear to me. Their similarity to ganglion-cells is not to be denied, though a connection with nerves could not be recognised. FHARYNGBAL GLAND-CELLS OP EARTHWORMS. 255 The matter is best left undecided at present." The projecting lobules on the dorsum of the pharynx Claparede called " ganglia of the previously described pharyngeal plexus." Vejdovsky's own account of the pharyngeal cells is not very clear, and is interpolated here and there amongst descriptions of the muscular and vascular apparatus of the pharynx,and of the occurrence and mechanism of its extrusion. Unlike Claparede, who recognised the identity of the cells of the lobules with those which penetrate inwards between the muscular strands (interpreting both as nervous), Vejdovsky considers them as distinct. Those which penetrate inwards he looks on simply as cellular elements of the coelomic fluid, which become attached to the pharyngeal muscles as to other organs; and he makes the rather surprising statement that " had Claparede compared these cells with those suspended in the coelomic fluid, he would certainly have recognised, them as the latter." The projecting lobules, on the other hand, are interpreted as mucous glands (Schleimdriisen) ; in vertical sections the glandular masses, contracting anteriorly to form long ducts, wind between the muscular bundles of the pharynx, and most probably empty their secretion into the pharyngeal cavity; these glands extend backwards far into cesophageal segments, and correspond to the septal or mucous glands of other Oligochseta. Vejdovsky also describes the ducts of the "septal glands" of Criodrilus as winding through the layer of muscular and vascular tissue on the dorsum of. the pharynx,, and the exceptionally large and numerous mucous glands of Dendrobsena rubida are said to consist each of a pear-shaped mass of cells.with large round nuclei and containing a substance which stains deeply in picrocarmine. •Vogt and Yung (10, 1888) describe irregularly dispersed cells between the muscular fibres on the dorsum of the pharynx (in Lumbricus agricola). These cells have ill- defined outlines, a granular protoplasm, and a clear spherical nucleus containing a nucleolus. The authors refer to •Claparede's interpretation of them as nerve cells; they 256 • • . j.- STEPHENSON. resemble, however,, the unicellular glands found in a corre- sponding position in other animals; and though the authors had not succeeded in discovering their ducts, they thought it not impossible that they secrete the viscid substance which the worm mixes with its food. Hesse (6, 1894) considers the pharyngeal cells of Oligo- chseta in general as belonging to the.epithelial layer; in the Naididse and Tubificidse the ventral end of each cell is pro- longed into a duct, which debouches between the lining epithelial cells of the pharyngeal cavity; the ducts of these cells are more drawn out in Lumbr.icus. Beddard (2, 1895) does not treat of the pharyngeal gland- cells of earthworms apart from the well-known "septal glands " of Enchytrasidse, etc. The septal glands in general, and so by implication the cells under consideration, appear to him to be simply epidermic glands which have been invaginated along with the storaodaeum, though their position causes him some doubt. The author who has examined these cellular aggregates in detail in the largest number of species, and who has given the most precise accounts of their supposed ductules and manner of discharge is Eisen (4, ft, 1895, 1896). In Phoenicodrilus taste the masses (called "salivary glands ") discharge through ducts which follow the muscle strands into the pharyngeal cavity j and it is probable that all the suprapharyngeal glands in Lumbricids open similarly and without any great variation as to detail: narrow ducts penetrate the pharyngeal epithelium, forming near the free surface small ovoid pockets for temporarily storing a small amount of the salivary secretion. These (suprapharyngeal) glands are connected posteriorly with the septal glands,— four pairs, superposed on several main longitudinal muscular bands which connect the pharyngeal glands with the body- wall in segment IX; their ducts., both wide and narrow, follow these muscles, so that the secretion- of the septal glands also is emptied into the pharynx. In Pontodrilus michaelseni the pharyngeal or salivary glands have a similar position, PHARYNGEAL GLAND-CELLS OP EARTHWORMS. 257 and are directly connected by means of duct3 with the epithelium of the pharynx; arrived at the pharyngeal epithelium the ducts branch out, sending numerous discharge tubes between the epithelial cells; these ductules are fre- quently, though not generally, branched while in the epithelial layer, and each ductule is furnished at the distal end with a small storage chamber of oblong form and con- siderably smaller than the nucleus of the epithelial cells. There are also in this species five pairs of septal glands, ventral to the oesophagus, and principally attached to blood- vessels, in segments V-IX, of similar structure ; a very thin duct runs backwards and upwards from the upper end of each towards the alimentary canal at its junction with the septum, " but I have some doubt about it emptying into the intestine, and it is much more probable that . these septal glands empty into the pharynx. None of my sections however show this to be the case." • The distribution of the septal glands in this species may be compared with what is found in Helodrilus (Bimastus) parvus (v. p. 23 post.). In Benhamia nan a Eisen states that the glands are evidently unicellular, and the fine ducts penetrate between the epithelial cells of the pharynx, the discharge pockets being almost globular; here and there the duct of a single glandular cell may be followed clear to the discharge pocket. " But to draw the conclusion . that all the pharyngeal and septal glands are unicellular is, I think, premature. In Pontodrilus, at least, there may be seen plainly numerous nuclei on the gland ducts, which of course indicates that we have here a fusion of several cells. ... In Pontodrilus the majority, and,.all the large glands, consisted of several cells, the respective ducts of which finally united into one. In Benhamia I could see no such union, and the single ducts could be followed with great facility to the outlets." In Benhamia nan a the septal glands, in segments IX, X, and XI, are very narrow and only one cell thick in the row. In B. palmicola the small septal glands are in'IX and X, lint 258 J. STEPHENSON. the author could not find that they were in any way connected with the pharyngeal system of glands. • The pharyngeal and septal glands of Aleodrilus keyesi are also described. Here it will be sufficient to call attention to the author's statements regarding the discharge of the gland-cells. The pharyngeal glands have discharge pockets which are much thicker than those seen in any other species; the septal glands are of the same nature as the pharyngeal, " but I have good reasons to believe that the glands in this species discharge into the tubular intestine.
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